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	<title>Biochemical Soul &#187; flatfish</title>
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		<title>Flatfish Eye Development &#8211; Video Update</title>
		<link>http://biochemicalsoul.com/2009/03/flatfish-eye-development-video-update/</link>
		<comments>http://biochemicalsoul.com/2009/03/flatfish-eye-development-video-update/#comments</comments>
		<pubDate>Sat, 07 Mar 2009 17:22:04 +0000</pubDate>
		<dc:creator>Irradiatus</dc:creator>
				<category><![CDATA[Biology]]></category>
		<category><![CDATA[Creatures]]></category>
		<category><![CDATA[Developmental Biology]]></category>
		<category><![CDATA[Fish]]></category>
		<category><![CDATA[Science]]></category>
		<category><![CDATA[development]]></category>
		<category><![CDATA[Evolution]]></category>
		<category><![CDATA[eye]]></category>
		<category><![CDATA[flatfish]]></category>
		<category><![CDATA[flounder]]></category>
		<category><![CDATA[Recapitulation Theory]]></category>

		<guid isPermaLink="false">http://biochemicalsoul.com/?p=1140</guid>
		<description><![CDATA[If you haven't read my piece on Flatfish Eyes &#38; Recapitulation Theory, you should check it out. For those of you who have read it, I updated it with the following AMAZING morph animations of flatfish development that I somehow missed before (much thanks to Adrian Thysse, FCD of Evolving Complexity for pointing these out [...]]]></description>
			<content:encoded><![CDATA[<p>If you haven't read my piece on <a href="../2009/02/2009/02/adaptation-of-the-week-flatfish-recapitulation/">Flatfish Eyes &amp; Recapitulation Theory</a>, you should check it out. For those of you who have read it, I updated it with the following AMAZING morph animations of flatfish development that I somehow missed before (much thanks to Adrian Thysse, FCD of <a href="http://evolvingwithdarwin.blogspot.com/" target="_blank">Evolving Complexity</a> for <a href="http://evolvingwithdarwin.blogspot.com/2008/07/hopeful-monster-flounders.html" target="_blank">pointing these out to me</a>).</p>
<p style="text-align: center;"><object width="400" height="302" data="http://vimeo.com/moogaloop.swf?clip_id=1309399&amp;server=vimeo.com&amp;show_title=1&amp;show_byline=0&amp;show_portrait=0&amp;color=&amp;fullscreen=1" type="application/x-shockwave-flash"><param name="allowfullscreen" value="true" /><param name="allowscriptaccess" value="always" /><param name="src" value="http://vimeo.com/moogaloop.swf?clip_id=1309399&amp;server=vimeo.com&amp;show_title=1&amp;show_byline=0&amp;show_portrait=0&amp;color=&amp;fullscreen=1" /></object><br />
<a href="http://vimeo.com/1309399">WANDERING EYES</a> from <a href="http://vimeo.com/sciencenews">Science News</a> on <a href="http://vimeo.com">Vimeo</a>.</p>
<p style="text-align: center;"><object width="400" height="283" data="http://vimeo.com/moogaloop.swf?clip_id=1309417&amp;server=vimeo.com&amp;show_title=1&amp;show_byline=0&amp;show_portrait=0&amp;color=&amp;fullscreen=1" type="application/x-shockwave-flash"><param name="allowfullscreen" value="true" /><param name="allowscriptaccess" value="always" /><param name="src" value="http://vimeo.com/moogaloop.swf?clip_id=1309417&amp;server=vimeo.com&amp;show_title=1&amp;show_byline=0&amp;show_portrait=0&amp;color=&amp;fullscreen=1" /></object><br />
<a href="http://vimeo.com/1309417">FROM FRY TO FISH</a> from <a href="http://vimeo.com/sciencenews">Science News</a> on <a href="http://vimeo.com">Vimeo</a>.</p>
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		<item>
		<title>Biochemical Sole</title>
		<link>http://biochemicalsoul.com/2009/03/biochemical-sole/</link>
		<comments>http://biochemicalsoul.com/2009/03/biochemical-sole/#comments</comments>
		<pubDate>Fri, 06 Mar 2009 00:53:44 +0000</pubDate>
		<dc:creator>Irradiatus</dc:creator>
				<category><![CDATA[Biochemicalsoul News]]></category>
		<category><![CDATA[Carnivals]]></category>
		<category><![CDATA[Funny]]></category>
		<category><![CDATA[Bedbugs]]></category>
		<category><![CDATA[Biochemical]]></category>
		<category><![CDATA[flatfish]]></category>
		<category><![CDATA[Malaria]]></category>
		<category><![CDATA[Sea Lice & Sunsets]]></category>
		<category><![CDATA[Sole]]></category>

		<guid isPermaLink="false">http://biochemicalsoul.com/?p=1130</guid>
		<description><![CDATA[The above image arrived in my email inbox last night from the preeminent coral reef expert and blogger extraordinaire Rick MacPherson of Malaria, Bedbugs, Sea Lice &#38; Sunsets. The subject line simply read "I'm just sayin..." The pun came from his recent Carnival of the Blue #22 in which he wrote of my post on [...]]]></description>
			<content:encoded><![CDATA[<div class="wp-caption aligncenter" style="width: 482px"><a href="http://coralnotesfromthefield.blogspot.com/2009/03/carnival-of-blue-22.html"><img title="Biochemical Sole" src="http://biochemicalsoul.com/images/sole.001.jpg" alt="In honor of the flatfishes..." width="472" height="241" /></a><p class="wp-caption-text">In honor of the flatfishes...</p></div>
<p>The above image arrived in my email inbox last night from the preeminent coral reef expert and blogger extraordinaire Rick MacPherson of <a href="http://coralnotesfromthefield.blogspot.com" target="_blank">Malaria, Bedbugs, Sea Lice &amp; Sunsets</a>.</p>
<p>The subject line simply read "I'm just sayin..."</p>
<p>The pun came from his recent <a href="http://coralnotesfromthefield.blogspot.com/2009/03/carnival-of-blue-22.html" target="_blank">Carnival of the Blue #22</a> in which he wrote of my post on <a href="../2009/02/adaptation-of-the-week-flatfish-recapitulation/">Flatfish Eyes &amp; Recapitulation Theory</a> :</p>
<blockquote><p><span style="font-weight: bold;">More Weird Fish Eyes</span><br />
If you still need some additional fish weirdness, Carnival of the Blue first-timer (hopefully long-timer) Daniel Brown of <span style="font-style: italic;"><a href="../">Biochemical Soul</a></span> dredges-up memories of 9AM Developmental Biology class as he explores <a href="../2009/02/adaptation-of-the-week-flatfish-recapitulation/">the evolution of flatfish eyes</a>. This post has it all... some ontogeny, some phylogeny, some eyeball migration. Perhaps with the right encouragement we can get Daniel to focus exclusively on the ocean and change his blog title to <span style="font-style: italic;">Biochemical Sole</span>?</p></blockquote>
<p>Rick's wit never ceases...</p>
<p>Go check him out to see what I mean.</p>
<p>I'm actually finding myself craving fish now.</p>
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		<title>Adaptation of the Week &#8211; Flatfish Eyes &amp; Recapitulation Theory</title>
		<link>http://biochemicalsoul.com/2009/02/adaptation-of-the-week-flatfish-recapitulation/</link>
		<comments>http://biochemicalsoul.com/2009/02/adaptation-of-the-week-flatfish-recapitulation/#comments</comments>
		<pubDate>Thu, 26 Feb 2009 00:17:45 +0000</pubDate>
		<dc:creator>Irradiatus</dc:creator>
				<category><![CDATA[Adaptation of the Week]]></category>
		<category><![CDATA[Biology]]></category>
		<category><![CDATA[Creatures]]></category>
		<category><![CDATA[Developmental Biology]]></category>
		<category><![CDATA[Evolution]]></category>
		<category><![CDATA[Fish]]></category>
		<category><![CDATA[Science]]></category>
		<category><![CDATA[development]]></category>
		<category><![CDATA[eye]]></category>
		<category><![CDATA[flatfish]]></category>
		<category><![CDATA[flounder]]></category>
		<category><![CDATA[Friedman]]></category>
		<category><![CDATA[Haeckel]]></category>
		<category><![CDATA[Recapitulation Theory]]></category>
		<category><![CDATA[Schreiber]]></category>

		<guid isPermaLink="false">http://biochemicalsoul.com/?p=1019</guid>
		<description><![CDATA[Most biologists at one time or another in their training have learned of the 19th century theory expounded upon by Ernst Haeckel called "Recapitulation Theory". The theory's thesis: "Ontogeny recapitulates phylogeny." Don't worry - it's not as complicated as the biological jargon might imply. The idea boils down to a simple one - one that [...]]]></description>
			<content:encoded><![CDATA[<p>Most biologists at one time or another in their training have learned of the 19th century theory expounded upon by <a href="http://en.wikipedia.org/wiki/Ernst_Haeckel" target="_blank">Ernst Haeckel</a> called "<a href="http://en.wikipedia.org/wiki/Recapitulation_theory" target="_blank">Recapitulation Theory</a>".</p>
<p>The theory's thesis: "Ontogeny recapitulates phylogeny." Don't worry - it's not as complicated as the biological jargon might imply.</p>
<div class="wp-caption alignright" style="width: 310px"><a href="http://en.wikipedia.org/wiki/Recapitulation_theory"><img title="Haeckel Drawings" src="http://biochemicalsoul.com/images/science/flatfish/Haeckel_drawings_small.jpg" alt="Ernst Haeckels Drawings (1892 Romane copy)" width="300" height="250" /></a><p class="wp-caption-text">Ernst Haeckel&#39;s Drawings (1892 Romane copy). Note: Haeckel oversimplified these drawings. I use them here as a simple illustration of the concept of developmental similarity.</p></div>
<p>The idea boils down to a simple one - one that seemed to make sense in light of the fact that the science of developmental biology had only just begun from a systematic standpoint.  The idea: if you watch an organism develop from an embryo to an adult, you can watch it slowly move through the evolutionary steps that had created it. That is: development repeats evolution.</p>
<p>So a human embryo would first look like a fish, then a reptile, then a mammal, and finally a human. Of course, we now know that in a literal sense, the theory is completely and utterly wrong. No stage of human development, or of any other organism, correlates with a discrete step in evolution.</p>
<p>We are never fish. (Though we <em>do </em>have embryonic tails).</p>
<p>However, that doesn't mean that there aren't kernels of truth to the idea, if applied loosely. Take the most famous and classic example: embryonic human gills. You may have heard yourself that humans have gills as embryos. Unfortunately this claim arises from misconception and incomplete understanding of developmental biology. Humans do not - ever - have gills. But as embryos we do have "pharyngeal arches." These are little bumps around what you might consider the neck area of a developing embryo (see Haeckel's drawings above). And these little mounds of tissue do in fact remarkably resemble similar mounds found in fish - mounds that in fish develop into gills (<strong>Note:</strong> Haeckel vastly oversimplified these drawings. I use them here as a simple illustration of the concept of developmental similarity. See: <a href="http://zygote.swarthmore.edu/evo5.html" target="_blank">http://zygote.swarthmore.edu/evo5.html</a>. Thanks <a href="http://pleion.blogspot.com/" target="_blank"><cite class="fn"></cite></a><a class="url" rel="external nofollow" href="http://pleion.blogspot.com/">Bjørn</a>!).</p>
<p>One of the amazing aspects of developmental biology that much of the public does not generally understand is that evolution does not occur by adding new organs, appendages, or tissues to adult animals (whether through gradual steps or not). Evolution works by slowly sculpting the early embryonic clay of an organism.</p>
<p>Fish evolved these gill pouches as embryos - pouches that could then be sculpted into gills. As evolution waltzed and hopped along at its geological pace, genetic mutations began to change how these little mounds were sculpted, such that now in humans, these arches are sculpted into various parts of the face and head. A genetic program was already in place to control the shaping of the pouch. All that natural selection did was slightly tweak that program. For example, instead of a group of cells moving one direction, they moved another. Instead of becoming blood vessel cells, they became cartilage or bone cells.</p>
<p>Thus, while we now understand that we are not witnessing evolution in miniature during development, we are seeing <em>pieces </em>of our evolutionary history - little remnants that remind us of our relationships to our ancestors and also help inform us on what morphogenetic processes underlie evolution.</p>
<p>Which brings us to our adaptation of the week: the freaky asymmetric eyes of the flatfish.</p>
<div class="wp-caption aligncenter" style="width: 510px"><a href="http://www.seawater.no/fauna/Fisk/flyndrefisk.htm"><img title="Flatfish" src="http://biochemicalsoul.com/images/science/flatfish/flatfish_small.jpg" alt="Flatfish (www.seawater.no)" width="500" height="374" /></a><p class="wp-caption-text">Flatfish (www.seawater.no)</p></div>
<p>Most people have probably seen a flounder - one member of the flatfishes. They have adapted to lie amongst the silty ocean bottom, hidden from predators and prey, flat on their sides. For a normal fish this might be maladaptive - they would constantly have one eye buried in the sand. Of course the negative of being one-eyed might be offset by being much more camouflaged and undetectable.</p>
<p>Luckily for the flounder, the eye that should be buried in the sand has moved around its forehead so that both eyes are on one side.</p>
<p>The flatfish eye served as one line of attack against natural selection back in the day - and Darwin himself didn't quite know how to answer the charges. Evolutionary gradualism would predict that through successive steps, the eye slowly moved upward toward the forehead and eventually to the other side of the face. But what advantage could a slightly moved eye give, if it still was on the wrong side? Alternatively, as Richard Goldschmidt postulated in the 1930s and 40s, perhaps a single monstrous freakfish was born with both eyes on one side, and this allowed it to lie flat without losing half its vision. It could have then survived and had lots of little freak fish babies of its own.</p>
<p>So how did the flatfish become the strange creature it is now? Let's first look at the developmental biology of the flatfish eye.</p>
<p style="text-align: center;"><object width="400" height="302" data="http://vimeo.com/moogaloop.swf?clip_id=1309399&amp;server=vimeo.com&amp;show_title=1&amp;show_byline=0&amp;show_portrait=0&amp;color=&amp;fullscreen=1" type="application/x-shockwave-flash"><param name="allowfullscreen" value="true" /><param name="allowscriptaccess" value="always" /><param name="src" value="http://vimeo.com/moogaloop.swf?clip_id=1309399&amp;server=vimeo.com&amp;show_title=1&amp;show_byline=0&amp;show_portrait=0&amp;color=&amp;fullscreen=1" /></object><br />
<a href="http://vimeo.com/1309399">WANDERING EYES</a> from <a href="http://vimeo.com/sciencenews">Science News</a> on <a href="http://vimeo.com">Vimeo</a>.</p>
<p style="text-align: center;"><object width="400" height="283" data="http://vimeo.com/moogaloop.swf?clip_id=1309417&amp;server=vimeo.com&amp;show_title=1&amp;show_byline=0&amp;show_portrait=0&amp;color=&amp;fullscreen=1" type="application/x-shockwave-flash"><param name="allowfullscreen" value="true" /><param name="allowscriptaccess" value="always" /><param name="src" value="http://vimeo.com/moogaloop.swf?clip_id=1309417&amp;server=vimeo.com&amp;show_title=1&amp;show_byline=0&amp;show_portrait=0&amp;color=&amp;fullscreen=1" /></object><br />
<a href="http://vimeo.com/1309417">FROM FRY TO FISH</a> from <a href="http://vimeo.com/sciencenews">Science News</a> on <a href="http://vimeo.com">Vimeo</a>.</p>
<div class="wp-caption alignright" style="width: 243px"><a href="http://www.ncbi.nlm.nih.gov/pubmed/16449556?ordinalpos=2&amp;itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DefaultReportPanel.Pubmed_RVDocSum"><img title="Flatfish development" src="http://biochemicalsoul.com/images/science/flatfish/metamorph_small.jpg" alt="Flatfish Metamorphosis" width="233" height="500" /></a><p class="wp-caption-text">Flatfish Metamorphosis (Schreiber 2006)</p></div>
<p>It's been know for quite some time that flatfish larvae look like perfectly normal, symmetrical, and upright fish. The picture to the right is from a study by Alexander Schreiber in the <a href="http://www.ncbi.nlm.nih.gov/pubmed/16449556?ordinalpos=2&amp;itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DefaultReportPanel.Pubmed_RVDocSum" target="_blank">Journal of Experimental Biology</a> from 2006. As you can see, at early stages the larvae are normal, but progressively tilt and become horizontal as one eye moves across the face. He also showed in this study that eye movement and flattening behavior occur independently during development - but that's a much longer story.</p>
<p>Alright, so one eye gradually moves across the skull during development. What about during evolution? Do we have any clues as to the steps involved? Well, as many biologists know, the fossil record has now answered the question for us.</p>
<p>In a well-known study that was published last summer in <a href="http://www.ncbi.nlm.nih.gov/pubmed/18615083?ordinalpos=1&amp;itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DefaultReportPanel.Pubmed_RVDocSum" target="_blank">Nature</a> and received much media attention, Matt Friedman showed findings from a series of fossils delineating a clear gradual evolution from symmetrical to asymmetrical flaltfishes. (For excellent in-depth coverage looking at this study and the debate over flatfish evolution, see one of my favorite science bloggers, Ed Yong at <a href="http://scienceblogs.com/notrocketscience/2008/07/missing_link_flatfish_has_eye_thats_moved_halfway_across_its.php" target="_blank">Not Exactly Rocket Science</a>, and also see the popular science writer Carl Zimmer at <a href="http://blogs.discovermagazine.com/loom/2008/07/09/dawn-of-the-picasso-fish/" target="_blank">The Loom</a>, and GrrlScientist at <a href="http://scienceblogs.com/grrlscientist/2008/07/the_mysterious_origin_of_the_w_1.php" target="_blank">Living the Scientific Life</a>).</p>
<p style="text-align: left;">
<div class="wp-caption aligncenter" style="width: 510px"><a href="http://www.ncbi.nlm.nih.gov/pubmed/18615083?ordinalpos=1&amp;itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DefaultReportPanel.Pubmed_RVDocSum"><img title="Flatfish Evolution" src="http://biochemicalsoul.com/images/science/flatfish/flatfishtree_small.jpg" alt="Flatfish Evolution" width="500" height="258" /></a><p class="wp-caption-text">Flatfish Evolution (Friedman 2008)</p></div>
<p>The evolution of the flatfish eye seems to mirror what we see during development. Thus, here we have a case of ontogeny appearing to recapitulate phylogeny quite wonderfully. There are many excellent examples of this throughout the biological world, though few that show such incredible similarity between the two processes of development and evolution. Nonetheless, this isn't <em>really </em>evolution we're watching during flatfish development - we're merely seeing how slight changes of the developmental programs are themselves responsible for the changes we see over time through evolution. Generally speaking, earlier developmental processes appear much more similar across varied species than later processes.</p>
<p>Development is in fact one of the primary constraints against evolution.</p>
<p>So while you were never a fish, you still showed remnants of fishy development during your own development. For it was these fishy developmental process that allowed the evolution of your own.</p>
<h2><strong>Scientific References:</strong></h2>
<ul>
<li>Schreiber AM. <a href="http://www.ncbi.nlm.nih.gov/pubmed/16449556?ordinalpos=2&amp;itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DefaultReportPanel.Pubmed_RVDocSum" target="_blank">Asymmetric craniofacial remodeling and lateralized behavior in larval flatfish</a>. <em>J Exp Biol.</em> 2006 Feb;209(Pt 4):610-21.</li>
<li>Friedman M. <a href="http://www.ncbi.nlm.nih.gov/pubmed/18615083?ordinalpos=1&amp;itool=EntrezSystem2.PEntrez.Pubmed.Pubmed_ResultsPanel.Pubmed_DefaultReportPanel.Pubmed_RVDocSum" target="_blank">The evolutionary origin of flatfish asymmetry</a>. <em><span class="journalname" title="Nature">Nature</span></em>. 2008 Jul 10;454(7201):209-12.</li>
</ul>
<h2><strong>Previous <a href="http://biochemicalsoul.com/category/adaptation-of-the-week/" target="_blank">Adaptations of the Week</a>:</strong></h2>
<ol>
<li><a href="http://biochemicalsoul.com/2009/01/adaptation-of-the-week-timber-rattlesnake-camouflage/" target="_blank">Timber Rattlesnake Camoflage</a></li>
<li><a href="http://biochemicalsoul.com/2009/02/adaptation-of-the-week-the-aye-ayes-freaky-finger-ive-been-cursed-by-an-aye-aye/" target="_blank">The Aye-Aye’s Freaky Finger (I’ve Been Cursed by an Aye-Aye!)</a></li>
</ol>
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